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Status, Ecology, Behaviour and Conservation of Raso Lark Alauda Razae

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Status, Ecology, Behaviour and Conservation of Raso Lark Alauda Razae Bird Conservation International (2003) 13:13–28. BirdLife International 2003 DOI: 10.1017/S0959270903003022 Printed in the United Kingdom Status, ecology, behaviour and conservation of Raso Lark Alauda razae PAUL F. DONALD, MARTA DE PONTE, MARIA J. PITTA GROZ and ROY TAYLOR Summary In October 2001, the world population of Raso Lark Alauda razae, confined to the uninhabited, arid islet of Raso in the Cape Verde Islands, was estimated at between 128 and 138 birds, of which 61–66% were males. The biased sex ratio was confirmed by standardized observations of randomly selected birds. The male-dominated sex ratio may result from differences in bill morphology and feeding methods. The males spent much time digging for the bulbs of the nutsedge Cyperus bulbosus, whereas females were seen to dig far less frequently. These differences may have been due to significant sexual differences in bill size. Both sexes also took a range of invertebrate prey, particularly lepidopteran larvae. Unpaired males showed differences in behaviour to paired males, particularly with respect to song flight length. Measurement of museum skins suggested significant seasonal changes in bill size, possibly related to changes in diet or feeding methods. Bill sizes in both sexes were highly variable, but significantly more variable in males. Nest predation, almost certainly by a near-endemic gecko, was extremely high, Mayfield estimates suggesting a nest survival rate of less than 5% between the onset of laying and the end of incubation. Cats and dogs were thought to be absent, although analysis of droppings shows that both have been present in the recent past. Analysis of historical data shows a strong correlation between population size and rainfall, and numbers of birds have fallen to extremely low levels during droughts. The conservation of the species is discussed in the light of these findings. Introduction Raso Lark Alauda razae is confined to the uninhabited 7 km2 island of Raso (16°37′N, 24°36′W) in the Cape Verde Islands. Discovered as recently as 1897 (Alexander 1898a), the species probably derived from Alauda-like stock in the Sahara during wetter glacial intervals (Hazevoet 1995), although its exact taxo- nomic affinities are unclear. It is possible that the species formerly had a larger range, as the present islands of Raso, Branco, Santa Luzia and Sa˜o Vicente were joined as a single island during glacial periods of lower sea levels (Hazevoet 1995), and sub-fossil bones found recently on Sa˜o Vicente may belong to this species. The current distribution may therefore be a relic of its former range (Bourne 1955, den Hartog 1990), although the species may have evolved in situ on Raso (Hazevoet 1995). Raso Lark is listed as Critically Endangered by IUCN (BirdLife International 2000) because of its very small distribution and small and fluctuating population. Since 1981, population estimates have ranged between 20 and 250 birds, although the reliability of counts is likely to be highly variable P. F. Donald et al. 14 (Ratcliffe et al. 1999). It is probably the least known bird species in the western Palaearctic, due largely to the inaccessibility of Raso and the absence of an orni- thological organization in the Cape Verde Islands. Most of what is known derives from the findings of ornithologists visiting the island for short periods, usually to study other species. Previous authors (e.g. Hazevoet 1995, Ratcliffe et al. 1999) have suggested that the population fluctuates in response to rainfall and that long-term desertification is a major threat. This is supported by the observation that birds are largely confined to small areas of green vegetation on the island’s flat plain (4 km2) and that breeding only seems to take place after rain (a common strategy in African larks), which falls largely in August to October (Alexander 1898b, Hazevoet 1995, Ratcliffe et al. 1999). As the Cape Verde Islands lie just outside the Intertropical Convergence Zone (ITCZ), periodic droughts are a feature of the climate of the archipelago (Hazevoet 1995), and Raso Lark has clearly survived droughts last- ing at least as long as the documented maximum of 18 years. Being relatively flat (maximum altitude 164 m) and having no permanent water source, water shortage on Raso is particularly severe and so the island has never been inhab- ited. During the 1968–1980 drought, numbers of Raso Larks may have fallen to less than 20 individuals (Ratcliffe et al. 1999). It has been suggested that nest predation may represent a further threat to this species (Castell 1999) and feral cats Felis catus have been present on the island in the recent past (Ratcliffe et al. 1999). The effects of predation may pose a particularly serious threat to the popu- lation when it has already been reduced to low levels by drought, so a critical assessment of all threats is necessary to formulate a conservation strategy for the species. This has hitherto been impossible with the little information available. This paper describes the results of an expedition to Raso in October 2001. The aims of the research were to assess the species’ status, diet, feeding methods and nesting success in order to understand the main threats faced by the species, and to collect documentary data to help clarify the species’ taxonomic affinities (Hazevoet 1989) and expand and update standard accounts (e.g. Cramp 1988). A specific aim was to locate and monitor nests to verify the assertion of Castell (1999) that nest predation rates may be high. Methods Visits were made to Raso on 30 September–4 October, 7–12 October, 16 October and 18 October 2001. The island was reached by chartered fishing boat from Tarrafal on the island of Sa˜o Nicolau, approximately 25 km to the east of Raso. Plant specimens were identified from Lobin (1982, 1986). Counts and assessments of sex ratio The whole of the flat plain of Raso (4 km2) was covered by four observers walk- ing parallel transects of 50 m length on October 1 and 3. A portable global posi- tioning system (GPS) was used to ensure full coverage and to avoid double counting. The exact location of all birds seen was recorded on the GPS to produce a distribution map. Males and females were distinguished in the counts based on bill size and shape and behaviour. It was not possible to cover the island’s Status, ecology, behaviour and conservation of Raso Lark 15 hilly northern side (3 km2) with parallel transects, so observers counted the small number of birds present there by visiting all patches of vegetation, following Ratcliffe et al.(1999). Initial results suggested an uneven sex ratio, possibly due to under-recording of females. Females may have been less conspicuous because they did not sing or because they may have been incubating eggs. Therefore, a standardized protocol was developed to determine the sex ratio more accurately. The GPS locations of 30 males were selected at random from those of all birds recorded during transects. An observer returned to this location and selected the male nearest to the point as the focal bird. Where no males were immediately obvious, the observer walked a spiral of 10 m intervals away from the point and the first male seen within 25 m of the observer selected as the focal bird. This was done to ensure that singing males were not selected in preference to males closer to the observer that were not singing. The focal bird was then watched for a period of one hour or until it was seen to be paired (in which case the time taken to ascertain this was recorded). The critical period of one hour was derived from observations of incubating females which showed that a male with an incubating female would encounter the female more than three times per hour and spend long periods with her (see Results). Therefore, a male observed for one hour without encountering a female could be safely assumed unpaired. The duration of song flights was also recorded to assess differences in territorial beha- viour that could also indicate a male’s status. Feeding and diet Observations were made of foraging behaviour of male and female birds to test previous assertions that the sexes feed in different ways. The focal bird was sexed on body size, bill size and shape, and behaviour (particularly song and crest-raising), and a maximum of 10 foraging actions recorded for each bird. A foraging action was defined as a successful attempt to gather food. These were recorded under one of four categories: digging, gleaning from vegetation, glean- ing from the ground and turning over stones. Significant differences in feeding methods between males and females were assessed using contingency table ana- lysis. Where possible, the food item was identified. Faecal samples were also collected from the song posts of a number of males, stored dry and analysed under a microscope to identify the main dietary components from undigested body parts. In order to determine whether any differences in feeding methods could be related to bill structure, an analysis was carried out to determine correlates of bill size. Bill measurements were taken from 45 museum skins (24 in the British Museum of Natural History, Tring, eight in the American Museum of Natural History, New York, five in the Muse´um National d’Histoire Naturelle, Paris and eight in the Centro de Zoologia, Instituto de Investigac¸a˜o Cientı´fica Tropical, Lisbon). Bill measurements taken were maximum depth and breadth, the length of the upper mandible from feathering on the forehead to bill tip and distance from the notch in the lower mandible to bill tip. Maximum bill length was mod- elled with general linear models (GLMs) in terms of sex, year of collection and month of collection (the last two to assess long-term or seasonal changes in bill size).
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